Cephalopod intelligence

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Two-thirds of an octopus's neurons are in the nerve cords of its tentacles. These are capable of complex reflex actions without input from the brain. Octopus tentacles.jpg
Two-thirds of an octopus's neurons are in the nerve cords of its tentacles. These are capable of complex reflex actions without input from the brain.

Cephalopod intelligence is a measure of the cognitive ability of the cephalopod class of molluscs.

Contents

Intelligence is generally defined as the process of acquiring, storing, retrieving, combining, comparing, and recontextualizing information and conceptual skills. [2] Though these criteria are difficult to measure in nonhuman animals, cephalopods are the most intelligent invertebrates. The study of cephalopod intelligence also has an important comparative aspect in the broader understanding of animal cognition because it relies on a nervous system fundamentally different from that of vertebrates. [3] In particular, the Coleoidea subclass (cuttlefish, squid, and octopuses) is thought to be the most intelligent invertebrates and an important example of advanced cognitive evolution in animals, though nautilus intelligence is also a subject of growing interest among zoologists. [4]

The scope of cephalopod intelligence and learning capability is controversial within the biological community, complicated by the inherent complexity of quantifying non-vertebrate intelligence. In spite of this, the existence of impressive spatial learning capacity, navigational abilities, and predatory techniques in cephalopods is widely acknowledged. [5] [6] Cephalopods have been compared to hypothetical intelligent extraterrestrials, due to their independently evolved mammal-like intelligence. [7]

Brain size and structure

Cephalopods have large, well-developed brains, [8] [9] [10] and their brain-to-body mass ratio is the largest among the invertebrates, falling between that of endothermic and ectothermic vertebrates. [11]

The nervous system of cephalopods is the most complex of all invertebrates. [10] [12] The giant nerve fibers of the cephalopod mantle have been widely used for many years as experimental material in neurophysiology; their large diameter (due to lack of myelination) makes them relatively easy to study compared with other animals. [13]

Behavior

Predation

A veined octopus eating a crab. Veined Octopus - Amphioctopus Marginatus eating a Crab.jpg
A veined octopus eating a crab.

Unlike most other molluscs, all cephalopods are active predators (with the possible exceptions of the bigfin squid and vampire squid). Their need to locate and capture their prey has likely been the driving evolutionary force behind the development of their intelligence. [14]

Crabs, the staple food source of most octopus species, present significant challenges with their powerful pincers and their potential to exhaust the cephalopod's respiration system from a prolonged pursuit. In the face of these challenges, octopuses will instead seek out lobster traps and steal the bait inside. They are also known to climb aboard fishing boats and hide in the containers that hold dead or dying crabs. [15] [16]

Captive cephalopods have also been known to climb out of their tanks, maneuver a distance of the lab floor, enter another aquarium to feed on the crabs, and return to their own aquariums. [17] [18] [19]

Communication

Although believed to not be the most social of animals, many cephalopods are in fact highly social creatures; when isolated from their own kind, some species have been observed shoaling with fish. [20]

Cephalopods are able to communicate visually using a diverse range of signals. To produce these signals, cephalopods can vary four types of communication elements: chromatic (skin coloration), skin texture (e.g. rough or smooth), posture, and locomotion. Changes in body appearance such as these are sometimes called polyphenism. [21] Some cephalopods are capable of rapid changes in skin colour and pattern through nervous control of chromatophores. [22] This ability almost certainly evolved primarily for camouflage, but squid use color, patterns, and flashing to communicate with each other in various courtship rituals. [21] Caribbean reef squid can even discriminate between recipients, sending one message using color patterns to a squid on their right, while they send another message to a squid on their left. [23] [24] Octopuses have been found to become more sociable when exposed to the psychoactive drug MDMA. [25]

The Humboldt squid shows extraordinary cooperation and communication in its hunting techniques. This is the first observation of cooperative hunting in invertebrates. [26]

It is believed that squids are slightly less intelligent than octopuses and cuttlefish; however, various species of squid are much more social and display greater social communications, etc., leading to some researchers concluding that squids are on par with dogs in terms of intelligence. [27]

Learning

A cuttlefish employing camouflage in its natural habitat. Camouflage cuttlefish 02.jpg
A cuttlefish employing camouflage in its natural habitat.

In laboratory experiments, octopuses can be readily trained to distinguish between different shapes and patterns, and one study concluded that octopuses are capable of using observational learning; [28] [29] however, this is disputed. [30] [31]

Octopuses have also been observed in what has been described as play: repeatedly releasing bottles or toys into a circular current in their aquariums and then catching them. [32]

Cephalopods can demonstrably benefit from environmental enrichment [33] indicating behavioral and neuronal plasticity not exhibited by many other invertebrates.

In a study on social learning, common octopuses (observers) were allowed to watch other octopuses (demonstrators) select one of two objects that differed only in color. Subsequently, the observers consistently selected the same object as did the demonstrators. [34]

Both octopuses and nautiluses are capable of vertebrate-like spatial learning. [35]

Tool use

A small coconut octopus (4-5 cm in diameter) using a nut shell and clam shell as shelter. Octopus shell.jpg
A small coconut octopus (4–5 cm in diameter) using a nut shell and clam shell as shelter.

The octopus has repeatedly been shown to exhibit flexibility in the use of tools.

At least four individuals of the veined octopus (Amphioctopus marginatus) have been observed retrieving discarded coconut shells, manipulating them, transporting them some distance, and then reassembling them for use as shelter. [36] It is surmised that the octopuses used bivalves for the same purpose before humans made coconut shells widely available on the sea floor. [37] [38] Other sea creatures construct homes in a similar manner; most hermit crabs use the discarded shells of other species for habitation, and some crabs place sea anemones on their carapaces to serve as camouflage. However, this behavior lacks the complexity of the octopus's fortress behavior, which involves picking up and carrying a tool for later use. (This argument remains contested by a number of biologists, who claim that the shells actually provide protection from bottom-dwelling predators in transport. [39] ) Octopuses have also been known to deliberately place stones, shells, and even bits of broken bottles to form walls that constrict their den openings. [40]

In laboratory studies, Octopus mercatoris, a small pygmy species of octopus, has been observed to block its lair using plastic Lego bricks. [41]

Smaller individuals of the common blanket octopus (Tremoctopus violaceus) hold the tentacles of the Portuguese man o' war (whose venom they are immune to), both as means of protection and as a method of capturing prey. [42]

Problem-solving ability

The highly sensitive suction cups and prehensile arms of octopuses, squid, and cuttlefish allow them to hold and manipulate objects. However, unlike vertebrates, the motor skills of octopuses do not seem to depend upon mapping their body within their brains, as the ability to organize complex movements is not thought to be linked to particular arms. [43]

Cephalopods can solve complex puzzles requiring pushing or pulling actions, and can also unscrew the lids of containers and open the latches on acrylic boxes in order to obtain the food inside. They can also remember solutions to puzzles and learn to solve the same puzzle presented in different configurations. [44]

Captive octopuses require stimulation or they will become lethargic; this typically takes the form of a variety of toys and puzzles. [45] At an aquarium in Coburg, Germany, an octopus named Otto was known to juggle his fellow tank-mates around, as well as throw rocks to smash the aquarium glass. On more than one occasion, Otto even caused short circuits by crawling out of his tank and shooting a jet of water at the overhead lamp. [46]

Additionally, cephalopods have been shown to have the capacity for future planning and reward processing after being tested with the Stanford marshmallow experiment. [47]

Protective legislation

An octopus in a zoo. Octopus.jpg
An octopus in a zoo.

Due to their intelligence, cephalopods are commonly protected by animal testing regulations that do not usually apply to invertebrates.

In the UK from 1993 to 2012, the common octopus ( Octopus vulgaris ) was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986. [48] Since 2022, all vertebrates, cephalopods, and decapods have been recognised as sentient by the Animal Welfare (Sentience) Act 2022.

Cephalopods are the only invertebrates protected under the 2010 European Union directive "on the protection of animals used for scientific purposes". [49]

In 2019, some scholars have argued for increased protections for cephalopods in the United States as well. [50]

See also

Related Research Articles

<span class="mw-page-title-main">Octopus</span> Soft-bodied eight-limbed order of molluscs

An octopus is a soft-bodied, eight-limbed mollusc of the order Octopoda. The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the center point of the eight limbs. The soft body can radically alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates.

<span class="mw-page-title-main">Squid</span> Superorder of cephalopod molluscs

A squid is a mollusc with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting these criteria. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.

<span class="mw-page-title-main">Cephalopod</span> Class of mollusks

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

<span class="mw-page-title-main">Vampire squid</span> Species of cephalopod

The vampire squid is a small cephalopod found throughout temperate and tropical oceans in extreme deep sea conditions. The vampire squid uses its bioluminescent organs and its unique oxygen metabolism to thrive in the parts of the ocean with the lowest concentrations of oxygen. It has two long retractile filaments, located between the first two pairs of arms on its dorsal side, which distinguish it from both octopuses and squids, and places it in its own order, Vampyromorphida, although its closest relatives are octopods. As a phylogenetic relict, it is the only known surviving member of its order.

<span class="mw-page-title-main">Animal cognition</span> Intelligence of non-human animals

Animal cognition encompasses the mental capacities of non-human animals including insect cognition. The study of animal conditioning and learning used in this field was developed from comparative psychology. It has also been strongly influenced by research in ethology, behavioral ecology, and evolutionary psychology; the alternative name cognitive ethology is sometimes used. Many behaviors associated with the term animal intelligence are also subsumed within animal cognition.

<span class="mw-page-title-main">Cephalization</span> Evolutionary trend of a head region developing

Cephalization is an evolutionary trend in animals that, over many generations, the special sense organs and nerve ganglia become concentrated towards the rostral end of the body where the mouth is located, often producing an enlarged head. This is associated with the animal's movement direction and bilateral symmetry, and cephalization of the nervous system led to the formation of a functional centralized brain in three groups of bilaterian animals, namely the arthropods, cephalopod molluscs, and vertebrates (craniates).

<span class="mw-page-title-main">Giant Pacific octopus</span> Species of cephalopod

The giant Pacific octopus, also known as the North Pacific giant octopus, is a large marine cephalopod belonging to the genus Enteroctopus. Its spatial distribution encompasses much of the coastal North Pacific, from Baja California state (Mexico), north along the United States' West Coast and British Columbia, Canada; across the northern Pacific to the Russian Far East, south to the East China Sea, the Yellow Sea, the Sea of Japan, Japan's Pacific east coast, and around the Korean Peninsula. It can be found from the intertidal zone down to 2,000 m (6,600 ft), and is best-adapted to colder, oxygen- and nutrient-rich waters. It is, arguably, the largest octopus species on earth and can often be found in aquariums and research facilities in addition to the ocean.

<span class="mw-page-title-main">Cephalopod limb</span> Limbs of cephalopod molluscs

All cephalopods possess flexible limbs extending from their heads and surrounding their beaks. These appendages, which function as muscular hydrostats, have been variously termed arms, legs or tentacles.

<span class="mw-page-title-main">Marine invertebrates</span> Marine animals without a vertebrate column

Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.

<span class="mw-page-title-main">Cuttlefish</span> Order of molluscs

Cuttlefish, or cuttles, are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of buoyancy.

<span class="mw-page-title-main">Cephalopod eye</span> Visual sensory organs of cephalopod molluscs

Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions. They have a camera-type eye which consists of an iris, a circular lens, vitreous cavity, pigment cells, and photoreceptor cells that translate light from the light-sensitive retina into nerve signals which travel along the optic nerve to the brain. For the past 140 years, the camera-type cephalopod eye has been compared with the vertebrate eye as an example of convergent evolution, where both types of organisms have independently evolved the camera-eye trait and both share similar functionality. Contention exists on whether this is truly convergent evolution or parallel evolution. Unlike the vertebrate camera eye, the cephalopods' form as invaginations of the body surface, and consequently the cornea lies over the top of the eye as opposed to being a structural part of the eye. Unlike the vertebrate eye, a cephalopod eye is focused through movement, much like the lens of a camera or telescope, rather than changing shape as the lens in the human eye does. The eye is approximately spherical, as is the lens, which is fully internal.

<span class="mw-page-title-main">Pain in animals</span> Overview about pain in animals

Pain negatively affects the health and welfare of animals. "Pain" is defined by the International Association for the Study of Pain as "an unpleasant sensory and emotional experience associated with actual or potential tissue damage, or described in terms of such damage." Only the animal experiencing the pain can know the pain's quality and intensity, and the degree of suffering. It is harder, if even possible, for an observer to know whether an emotional experience has occurred, especially if the sufferer cannot communicate. Therefore, this concept is often excluded in definitions of pain in animals, such as that provided by Zimmerman: "an aversive sensory experience caused by actual or potential injury that elicits protective motor and vegetative reactions, results in learned avoidance and may modify species-specific behaviour, including social behaviour." Nonhuman animals cannot report their feelings to language-using humans in the same manner as human communication, but observation of their behaviour provides a reasonable indication as to the extent of their pain. Just as with doctors and medics who sometimes share no common language with their patients, the indicators of pain can still be understood.

<span class="mw-page-title-main">Pain in crustaceans</span> Ethical debate

There is a scientific debate which questions whether crustaceans experience pain. It is a complex mental state, with a distinct perceptual quality but also associated with suffering, which is an emotional state. Because of this complexity, the presence of pain in an animal, or another human for that matter, cannot be determined unambiguously using observational methods, but the conclusion that animals experience pain is often inferred on the basis of likely presence of phenomenal consciousness which is deduced from comparative brain physiology as well as physical and behavioural reactions.

<span class="mw-page-title-main">Underwater camouflage</span> Camouflage in water, mainly by transparency, reflection, counter-illumination

Underwater camouflage is the set of methods of achieving crypsis—avoidance of observation—that allows otherwise visible aquatic organisms to remain unnoticed by other organisms such as predators or prey.

<span class="mw-page-title-main">Pain in invertebrates</span> Contentious issue

Pain in invertebrates is a contentious issue. Although there are numerous definitions of pain, almost all involve two key components. First, nociception is required. This is the ability to detect noxious stimuli which evokes a reflex response that moves the entire animal, or the affected part of its body, away from the source of the stimulus. The concept of nociception does not necessarily imply any adverse, subjective feeling; it is a reflex action. The second component is the experience of "pain" itself, or suffering—i.e., the internal, emotional interpretation of the nociceptive experience. Pain is therefore a private, emotional experience. Pain cannot be directly measured in other animals, including other humans; responses to putatively painful stimuli can be measured, but not the experience itself. To address this problem when assessing the capacity of other species to experience pain, argument-by-analogy is used. This is based on the principle that if a non-human animal's responses to stimuli are similar to those of humans, it is likely to have had an analogous experience. It has been argued that if a pin is stuck in a chimpanzee's finger and they rapidly withdraw their hand, then argument-by-analogy implies that like humans, they felt pain. It has been questioned why the inference does not then follow that a cockroach experiences pain when it writhes after being stuck with a pin. This argument-by-analogy approach to the concept of pain in invertebrates has been followed by others.

<span class="mw-page-title-main">Deimatic behaviour</span> Bluffing display of an animal used to startle or scare a predator

Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey animal an opportunity to escape. The term deimatic or dymantic originates from the Greek δειματόω (deimatóo), meaning "to frighten".

<span class="mw-page-title-main">Pain in cephalopods</span> Contentious issue

Pain in cephalopods is a contentious issue. Pain is a complex mental state, with a distinct perceptual quality but also associated with suffering, which is an emotional state. Because of this complexity, the presence of pain in non-human animals, or another human for that matter, cannot be determined unambiguously using observational methods, but the conclusion that animals experience pain is often inferred on the basis of likely presence of phenomenal consciousness which is deduced from comparative brain physiology as well as physical and behavioural reactions.

<i>Other Minds: The Octopus, the Sea, and the Deep Origins of Consciousness</i> 2016 book on evolution of consciousness by Peter Godfrey-Smith

Other Minds is a 2016 bestseller by Peter Godfrey-Smith on the evolution and nature of consciousness. It compares the situation in cephalopods, especially octopuses and cuttlefish, with that in mammals and birds. Complex active bodies that enable and perhaps require a measure of intelligence have evolved three times, in arthropods, cephalopods, and vertebrates. The book reflects on the nature of cephalopod intelligence in particular, constrained by their short lifespan, and embodied in large part in their partly autonomous arms which contain more nerve cells than their brains.

<span class="mw-page-title-main">Dwarf cuttlefish</span> Species of cuttlefish

The dwarf cuttlefish (Sepia bandensis), also known as the stumpy-spined cuttlefish, is a species of cuttlefish native to the shallow coastal waters of the Central Indo-Pacific. The holotype of the species was collected from Banda Neira, Indonesia. It is common in coral reef and sandy coast habitats, usually in association with sea cucumbers and sea stars. Sepia baxteri and Sepia bartletti are possible synonyms.

Octopus bocki is a species of octopus, which has been located near south Pacific islands such as Fiji, the Philippines, and Moorea and can be found hiding in coral rubble. They can also be referred to as the Bock's pygmy octopus. They are nocturnal and use camouflage as their primary defense against predators as well as to ambush their prey. Their typical prey are crustaceans, crabs, shrimp, and small fish and they can grow to be up to 10cm in size.

References

  1. Yekutieli, Y.; Sagiv-Zohar, R.; Aharonov, R.; Engel, Y.; Hochner, B.; Flash, T. (2005). "Dynamic model of the octopus arm. I. Biomechanics of the octopus reaching movement". Journal of Neurophysiology. 94 (2): 1443–1458. doi:10.1152/jn.00684.2004. PMID   15829594. S2CID   14711055.
  2. Humphreys, Lloyd G. (April–June 1979). "The construct of general intelligence" (PDF). Intelligence (editorial). 3 (2): 105–120. doi:10.1016/0160-2896(79)90009-6. ISSN   0160-2896. Archived (PDF) from the original on 12 August 2017. Retrieved 13 December 2020.
  3. "Cephalopod intelligence" Archived 2020-03-21 at the Wayback Machine in The Encyclopedia of Astrobiology, Astronomy, and Spaceflight.
  4. Crook, Robyn & Basil, Jennifer (2008). "A biphasic memory curve in the chambered nautilus, Nautilus pompilius L. (Cephalopoda: Nautiloidea)" (PDF). Journal of Experimental Biology . 211 (12): 1992–1998. doi: 10.1242/jeb.018531 . PMID   18515730. Archived (PDF) from the original on 4 November 2018. Retrieved 13 December 2020.
  5. Hunt, Elle (28 March 2017). "Alien intelligence: the extraordinary minds of octopuses and other cephalopods". The Guardian . Archived from the original on 18 April 2020.
  6. Bilefsky, Dan (April 13, 2016). "Inky the Octopus Escapes From a New Zealand Aquarium" . The New York Times . Archived from the original on 16 April 2020. Retrieved 24 April 2016.
  7. Baer, Drake (20 December 2016). "Octopuses Are 'the Closest We Will Come to Meeting an Intelligent Alien'". Science of Us. Retrieved 26 April 2017.
  8. Tricarico, Elena; Amodio, Piero; Ponte, Giovanna; Fiorito, Graziano (2014). "Cognition and recognition in the cephalopod mollusc Octopus vulgaris: coordinating interaction with environment and conspecifics". In Witzany, Guenther (ed.). Biocommunication of Animals. Springer. pp. 337–349. doi:10.1007/978-94-007-7414-8_19. ISBN   978-94-007-7413-1. LCCN   2019748877.
  9. Chung, Wen-Sung; Kurniawan, Nyoman D.; Marshall, N. Justin (2020). "Toward an MRI-Based Mesoscale Connectome of the Squid Brain". iScience. 23 (1): 100816. Bibcode:2020iSci...23j0816C. doi: 10.1016/j.isci.2019.100816 . ISSN   2589-0042. PMC   6974791 . PMID   31972515.
  10. 1 2 Chung, Wen-Sung; Kurniawan, Nyoman D.; Marshall, N. Justin (2021-11-18). "Comparative brain structure and visual processing in octopus from different habitats". Current Biology. 32 (1): 97–110.e4. doi: 10.1016/j.cub.2021.10.070 . ISSN   0960-9822. PMID   34798049. S2CID   244398601.
  11. Nixon, Marion; Young, John Z. (4 September 2003). The Brains and Lives of Cephalopods. Oxford University Press (published November 6, 2003). ISBN   978-0198527619. LCCN   2002041659.
  12. Budelmann, B. U. (1995). "The cephalopod nervous system: What evolution has made of the molluscan design". In Breidbach, O.; Kutsch, W. (eds.). The nervous systems of invertebrates: An evolutionary and comparative approach. Birkhäuser. ISBN   978-3-7643-5076-5. LCCN   94035125.
  13. Tasaki, I.; Takenaka, T. (October 1963). "Resting and action potential of squid giant axons intracellularly perfused with sodium-rich solutions" (PDF). Proceedings of the National Academy of Sciences of the United States of America . 50 (4): 619–626. Bibcode:1963PNAS...50..619T. doi: 10.1073/pnas.50.4.619 . PMC   221236 . PMID   14077488. Archived (PDF) from the original on 11 August 2018. Retrieved 13 December 2020.
  14. Villanueva, Roger; Perricone, Valentina; Fiorito, Graziano (2017-08-17). "Cephalopods as Predators: A Short Journey among Behavioral Flexibilities, Adaptions, and Feeding Habits". Frontiers in Physiology. 8: 598. doi: 10.3389/fphys.2017.00598 . ISSN   1664-042X. PMC   5563153 . PMID   28861006.
  15. Cousteau, Jacques Yves (1978). Octopus and Squid: The Soft Intelligence
  16. "Giant Octopus – Mighty but Secretive Denizen of the Deep". Smithsonian National Zoological Park. 2 January 2008. Archived from the original on 25 August 2012. Retrieved 4 February 2014.
  17. Wood, J. B; Anderson, R. C (2004). "Interspecific Evaluation of Octopus Escape Behavior" (PDF). Journal of Applied Animal Welfare Science. 7 (2): 95–106. doi:10.1207/s15327604jaws0702_2. PMID   15234886. S2CID   16639444 . Retrieved 11 September 2015.
  18. Lee, Henry (1875). "V: The octopus out of water". Aquarium Notes – The Octopus; or, the "devil-fish" of fiction and of fact. London: Chapman and Hall. pp. 38–39. OCLC   1544491 . Retrieved 11 September 2015. The marauding rascal had occasionally issued from the water in his tank, and clambered up the rocks, and over the wall into the next one; there he had helped himself to a young lump-fish, and, having devoured it, returned demurely to his own quarters by the same route, with well-filled stomach and contented mind.
  19. Roy, Eleanor Ainge (14 April 2016). "The great escape: Inky the octopus legs it to freedom from aquarium". The Guardian (Australia).
  20. Packard, A. (1972). "Cephalopods and fish: The limits of convergence". Biological Reviews. 47 (2): 241–307. doi:10.1111/j.1469-185X.1972.tb00975.x. S2CID   85088231.
  21. 1 2 Brown, C.; Garwood, M. P.; Williamson, J.E. (2012). "It pays to cheat: Tactical deception in a cephalopod social signalling system". Biology Letters. 8 (5): 729–732. doi:10.1098/rsbl.2012.0435. PMC   3440998 . PMID   22764112.
  22. Cloney, R.A.; Florey, E. (1968). "Ultrastructure of cephalopod chromatophore organs". Z. Zellforsch. Mikrosk. Anat. 89 (2): 250–280. doi:10.1007/BF00347297. PMID   5700268. S2CID   26566732.
  23. "Sepioteuthis sepioidea, Caribbean Reef squid". The Cephalopod Page. Retrieved 20 January 2010.
  24. Byrne, R.A.; Griebel, U.; Wood, J.B.; Mather, J.A. (2003). "Squids say it with skin: A graphic model for skin displays in Caribbean Reef Squid". Berliner Geowissenschaftliche Abhandlungen. 3: 29–35.
  25. Nuwer, Rachel. "Rolling under the Sea: Scientists Gave Octopuses Ecstasy to Study Social Behavior". Scientific American.
  26. Zimmermann, Tim (July 2006). "Behold the Humboldt squid". Outside Magazine.
  27. "Are squids as smart as dogs?". www.medicalnewstoday.com. 2020-02-10. Retrieved 2021-06-07.
  28. Fiorito, Graziano; Scotto, Pietro (24 April 1992). "Observational Learning in Octopus vulgaris". Science. 256 (5056): 545–547. Bibcode:1992Sci...256..545F. doi:10.1126/science.256.5056.545. PMID   17787951. S2CID   29444311 . Retrieved 18 February 2015.
  29. "Octopus intelligence: Jar opening". BBC News. 25 February 2003. Retrieved 4 February 2014.
  30. Hamilton, Garry (7 June 1997). "What is this octopus thinking?". New Scientist. No. 2085. pp. 30–35. Retrieved 18 February 2015.
  31. Stewart, Doug (1997). "Armed but not dangerous: Is the octopus really the invertebrate intellect of the sea". National Wildlife. 35 (2).
  32. Mather, J. A.; Anderson, R. C. (1998). Wood, J. B. (ed.). "What behavior can we expect of octopuses?". The Cephalopod Page.
  33. Mather, J.A., Anderson, R.C. and Wood, J.B. (2010). Octopus: The Ocean's Intelligent Invertebrate. Timber Press.{{cite book}}: CS1 maint: multiple names: authors list (link)
  34. Fiorito, G. & Scotto, P. (1992). "Observational learning in Octopus vulgaris". Science. 256 (5056): 545–547. Bibcode:1992Sci...256..545F. doi:10.1126/science.256.5056.545. PMID   17787951. S2CID   29444311.
  35. Crook, R.J. & Walters, E.T. (2011). "Nociceptive behavior and physiology of molluscs: animal welfare implications". ILAR Journal. 52 (2): 185–195. doi: 10.1093/ilar.52.2.185 . PMID   21709311.
  36. Finn, Julian K.; Tregenza, Tom; Norman, Mark D. (15 December 2009). "Defensive tool use in a coconut-carrying octopus" (PDF). Current Biology . 19 (23): R1069–R1070. doi: 10.1016/j.cub.2009.10.052 . PMID   20064403. Archived (PDF) from the original on 11 August 2017 via Occidental College.
  37. Morelle, Rebecca (14 December 2009). "Octopus snatches coconut and runs". BBC News . Archived from the original on 31 May 2020. Retrieved 20 January 2010.
  38. "Coconut shelter: evidence of tool use by octopuses | EduTube Educational Videos". Edutube.org. 2009-12-14. Archived from the original on 2013-10-24. Retrieved 2010-01-20.
  39. Octopus tool use on YouTube published January 26, 2010 New Scientist
  40. "Simple tool use in owls and cephalopods". Map Of Life. 2010. Retrieved July 23, 2013.
  41. Oinuma, Colleen, (14 April 2008). "Octopus mercatoris response behavior to novel objects in a laboratory setting: Evidence of play and tool use behavior?" In Octopus Tool Use and Play Behavior
  42. Jones, Everet C. (22 February 1963). "Tremoctopus violaceus uses Physalia tentacles as weapons". Science . 139 (3556): 764–766. Bibcode:1963Sci...139..764J. doi:10.1126/science.139.3556.764. JSTOR   1710225. PMID   17829125. S2CID   40186769.
  43. Zullo, Letizia; Sumbre, German; Agnisola, Claudio; Flash, Tamar; Hochner, Binyamin (17 September 2009). "Nonsomatotopic organization of the higher motor centers in octopus" (PDF). Current Biology . 19 (19): 1632–6. doi: 10.1016/j.cub.2009.07.067 . PMID   19765993. Archived (PDF) from the original on 9 July 2020. Retrieved 13 December 2020.
  44. Richter, Jonas N.; Hochner, Binyamin; Kuba, Michael J. (2016-03-22). "Pull or Push? Octopuses Solve a Puzzle Problem". PLOS ONE. 11 (3): e0152048. Bibcode:2016PLoSO..1152048R. doi: 10.1371/journal.pone.0152048 . ISSN   1932-6203. PMC   4803207 . PMID   27003439.
  45. "Captive Octopuses Need Intellectual Stimulation Or Else They Get Bored". curiosity.com. Retrieved 2018-11-19.[ permanent dead link ]
  46. "Otto the octopus wreaks havoc" . The Telegraph . 31 October 2008. Archived from the original on 24 June 2011.
  47. Starr, Michelle (3 March 2021). "A Cephalopod Has Passed a Cognitive Test Designed For Human Children". ScienceAlert. Retrieved 2021-03-03.
  48. "The Animals (Scientific Procedures) Act (Amendment) Order 1993". The National Archives. Retrieved 18 February 2015.
  49. "DIRECTIVE 2010/63/EU OF THE EUROPEAN PARLIAMENT AND OF THE COUNCIL". Official Journal of the European Union. Article 1, 3(b). Retrieved 18 February 2015.
  50. Zabel, Joseph (Spring 2019). "Legislators Need to Develop a Backbone for Animals that Lack One: Including Cephalopods in the Animal Welfare Act". Journal of Animal and Environmental Law. University of Louisville School of Law. 10 (2): 1.

Further reading